Frederick Hotchkiss, PhD

annotations to published articles

Hotchkiss, F.H.C. 2012. Growth zones and extraxial-axial skeletal homologies in Asteroidea. Proceedings of the Biological Society of Washington 125(2):106-121. Also given as a poster presentation titled " Revised extraxial-axial homologies in Asteroidea ", at the 14th International Echinoderms Conference, August 2012, Brussels, Belgium. MPRI contribution No. 6. Further to gusset plate terminology on p. 108: As well explained by Branstrator (1975), gusset plates increase the girth and the internal volume of the rays. He described a progression of development of intermarginal and dorsolateral gusset plates in Ordovician Asteroidea from not present in Hudsonasteridae, to present in Mesopalaeasteridae, and to proliferated presence in Promopalaeasteridae. Likewise a proliferated presence of dorsolaterals in Urasterellidae. Further to the list of families of mentioned genera of asteroids (materials section, p. 109): add Asteriidae: Leptasterias ochotensis similispinus. Further to the mention of Pteraster on p. 109 and p. 117, this may be a misidentification because the cited figure [R. Mooi & B. David 2000. What a new model of skeletal homologies tells us about asteroid evolution. American Zoologist 40(3):326-339, their figure 2 on page 330, captioned “Morphology of juvenile Pteraster? CASIZ 9175] shows and labels both inframarginal and supramarginal plates. We know from Villier, L., D.B. Blake, J.W.M. Jagt and M. Kutscher [2004. A preliminary phylogeny of the Pterasteridae (Echinodermata, Asteroidea) and the first fossil record: Late Cretaceous of Germany and Belgium. Paläontologische Zeitschrift 78(2):281-299, see p. 282] that Pteraster lacks marginals. Postscript to scope p. 109: Beyond the stated scope but of interest is that Mooi & David identify the primary central plate of the asteroid disk as vestigial imperforate extraxial body wall. Likewise of interest is that in ophiuroids the disk primary plates do not regenerate, whereas the radial shields, genital bars, etc. do regenerate; so fundamental differences as noted by Hendler (1978). Postscript to p. 109 OPR topic in materials and methods: Concerning echinoids, Mooi et al. (1994:88) and Mooi & David (1997:308) applied the OPR to a teratological specimen of Strongylocentrotus with an altered apical system (specimen of Jackson 1912:37) as a corroboration; likewise they applied the OPR to interpret the extreme morphology of the holasteroid apical system in Plexechinus cinctus as a corroboration. Their method of corroboration of how the OPR operates in echinoids nicely parallels the examination herein of arm stump specimens and especially Zirpolo’s (1928) undersurface branch arm specimen to reveal how the OPR operates in asteroids. Further to p. 115 discussion: Based on teratological specimens (FHCH 1979, p. 150), Tiedemann's bodies are present or absent as a pair (as a unit), one on each side of the ambulacral axis, conforming with identification of asteroid OPR growth zones. Concerning possibilities for future research, it is conceivable that the place of intra-disk autotomy discovered in Luidia clathrata by Schram & McClintock (2009. Rapid regeneration of the body wall of the aboral central disc in the sea star Luidia clathrata. Gulf of Mexico Science 27(2):123-124) is related to the boundary between extraxial and axial body wall in this species. Also deserving mention is the idea of Mooi & David (1997:329) that there may be "deep-seated ultrastructural differences in the stereom organization" of extraxial versus axial skeleton that could be "exploited to determine affinities of individual stereom elements, independent of topological position". Further to Fig. 5, it is self evident that the development of superomarginals behind the terminal plates continues to wall off the abactinal surface from the inferomarginals and from all of the subambital plates of the actinal surface. Therefore neither the inferomarginal plates nor any of the subambital plates can have anything to do with the abactinal surface, meaning that they cannot be extraxial plates.

Hotchkiss, F.H.C. and J.K. Keesing. 2012. An arm-stump specimen of Archaster angulatus Muller & Troschel, 1842 (Echinodermata: Asteroidea) [abstract]. Gulf of Mexico Science 29(2):144-145 [cover says 2011, but actual is June 2012]. Poster presentation prepared for the 6th North American Echinoderm Conference, Rosario Beach Marine Laboratory, Anacortes, WA, USA, August 14-19, 2011. MPRI contribution No. 5.

Hotchkiss, F.H.C. and A. Glass. 2010 and 2012. Observations on Onychaster (Ophiuroidea: Onychasteridae) (Famennian - Visean Age. Poster presentation prepared for the 7th European Conference on Echinoderms. MPRI contribution No. 4. Abstract published 2010: pp. 52-53 in M. Reich, J. Reitner, V. Roden, and B. Thuy (eds.), Echinoderm Research 2010, 7th European Conference on Echinoderms, Göttingen, October 2-9, 2010. Abstract volume and field guide to excursions. Universitätsdrucke Göttingen. Full paper published 2012 in Zoosymposia 7:121-138. Further to "Onychaster classified as a euryalid" in Table 1 on page 123, add Lyman, 1882, Challenger Report on the Ophiuroidea, see page 328: "It is plain that simple armed Astrophytons began as low as the coal; for Onychaster flexilis, Meek and Worthen, evidently belongs in this group." Further to "Onychaster classified as non-euryalid" in Table 1 on page 123, add MacBride, 1906, Echinodermata chapter in The Cambridge Natural History, vol. 1, 623 pp., MacMillan, London, see page 502: Onychaster "is a representative of the Streptophiurae". Ditto as non-euryalid add Sedgwick, 1909, A student's textbook of zoology, Vol. III (Chapter III Echinodermata; p. 206, Onychasteridae as Streptophiurae), Swan Sonnenschein and Co. Ltd, London, and MacMillan Co., NY, 905 pp. Further to the specimen data on pp. 124-125 I want to add that all of the ALABAMA specimens were collected by Richard Keyes, Huntsville, AL. Further to p. 131 comparison with functioning of cardinal teeth of pelecypod hinge: see p. 413 on Class Pelecypoda by John Pojeta, Jr., in textbook "Fossil Invertebrates", Blackwell Scientific Publications, editors Boardman, Cheetham, and Rowell, 1987.

Hotchkiss, F.H.C. 2008 - 2009. Pattern formation in starfish: arm stumps, regeneration models and evolution. MPRI contribution No. 3. Poster presentation prepared for the Fifth North American Echinoderm Conference, Melbourne, Florida, July 2008, and also the 4th Workshop of German and Austrian Echinoderm Research, Vienna, October 2008. Poster presented in 2009 at the 13th International Echinoderms Conference, Hobart, Tasmania; at the Western Interior Paleontological Society's Founders' Symposium; and at the 14th Annual Cape Cod Natural History Conference. Link to Vienna abstracts. 2009. Hotchkiss, F. H. C. Arm stumps and regeneration models in Asteroidea (Echinodermata). Proceedings of the Biological Society of Washington 122(3):342-354. Other published/unpublished mentions of ray stumps that do not regenerate : Ruedemann (1916:61) described arm stumps in Hallaster forbesi, but Spencer (Monograph p. 298) showed that the arms are flexed aborally giving only a false appearance of being stumps. The abruptly terminating arms of Cholaster peculiaris, which is known from a single specimen, are also the product of arms bent 180 degrees aborally onto themselves (FHCH unpublished). After the 2009 Hobart IEC meeting John Lawrence and John Keesing found off Perth, Australia, an Archaster with an arm stump that healed and did not regenerate, similar to the specimens reported in the poster. Sumrall, Brett & McKinney (2009 J. Paleont. 83:794-803, see p. 801 and fig. 7) describe a Multiplexidiscus mckenziensis edrioasterd specimen in which the B ambulacrum failed to fully develop: ray B developed instead as a blunt hemicircular ambulacral terminus. Additional examples of regenerating Ordovician starfish: Urasterella sp., ROM 53376, Middle Ordovician, Bobcaygeon Formation, near Lake Simcoe, Ontario, Canada (personal communication, David Rudkin, 4/15/2008). Cnemidactis girvanensis (Schuchert), counterpart specimens BMNH E 52484a,b; Upper Ordovician (Ashgillian), Thraive Glen, Ayreshire, Scotland; figured by W.K. Spencer (1918:161 and pl. XII fig. 5); Owen (1965:565 curatorial notes). Phyrtosaster casteri with disproportionately small ossicles at arm tip suggesting arm tip loss and subsequent regeneration (Blake 2007:1483, Fig. 3.7 caption). Chandebois (1973. Acta Biotheoretica 22(1):2-33; see pp. 16-18) gives more examples of wound healing without regeneration. (p. 349) First, in marine planarians “a blastema never appears when the cut surface contracts so that its two dorsal half-edges fuse together, and likewise the two ventral edges (Chandebois 1957a).” Planarian wound healing that joins dorsal tissue with ventral counterparts forms a blastema. Second (p. 18), reporting that Coulomb (1972) observed in the lumbricid oligochaete Eiseniella tetraedra that “a blastema appears if dorsal epidermis meets ventral epidermis, but it never forms when the two lateral halves of the wound join.” A seamless transition (p. 349) from regenerative growth to normal growth is supported by King's (1898:354) observations on sections through the arm tips of regenerating and normal arms. Indeterminate growth (p. 349) in the Asteriidae is supported by Verill's statement (1914 Harriman Alaska report, p. 31) that size and age have no definite limit. Indeterminate growth is not excluded in Acanthaster [Stump & Lucas. 1999. Age estimation and patterns of growth in Acanthaster planci: a reply to Souter et al. (1997). Marine and Freshwater Research 50(1):71-72]. Individuals of the echinoid Strongylocentrotus franciscanus continue to grow throughout life [T.A. Ebert & J.R. Southon. 2003. Red sea urchins (Strongylocentrotus fransciscanus) can live over 100 years: confirmation with A-bomb 14carbon. Fish. Bull. 101(4):915-922]. Indeterminate growth in asteroids and echinoids discussed in K.P. Sebens [1987. The ecology of indeterminate growth in animals. Ann. Rev. Ecol. Syst. 18:371-407]. Indeterminate growth in Pisaster ochraceus discussed by Carlos Robles [2013. see Chapt. 16 in book Starfish -- biology and ecology of the Asteroidea, J.M. Lawrence, ed., Johns Hopkins Univ. Press] Previous research/comments on symmetric versus asymmetric positional information in the starfish arm: Lovén’s rule and Batesonian duplications of appendages imply asymmetrical positional information. David, Mooi & Telford (Echinoderm Research 1995: 163): “In groups such as Asteroidea, in which the paired plates of the ambulacral columns are directly opposite and do not alternate, it should still be possible to see from early ontogenetic stages in which columns the first plate is laid down, thereby testing the existence of Lovén’s rule even in this group.” Hotchkiss (1998: 207) emphasized that no case of Batesonian duplication of asteroid rays has ever been reported, and he rejected the implications of this negative finding. The implied expectation of these researchers was that positional information in the starfish ray is asymmetric. Impact on the rays-as-appendages hypothesis: Symmetrical positional information in the starfish ray is not consistent with the rays-as-appendages model which explicitly invokes asymmetrical positional information. Either the rays-as-appendages hypothesis should be rejected, or we need some sort of hypothesis-saving explanation. I opt for the latter because the fossil record indicates that alternating ambulacral plates and Loven’s law are plesiomorphic, and these features imply asymmetric positional information. Somehow, the starfish arm has changed from an original asymmetric condition to a symmetric condition. Other examples of failure to regenerate following L-R bilaterally symmetrical wound closure and stump formation occur in planarians (headless planarians): Child, C.M. 1911. Experimental control of morphogenesis in the regulation of Planaria. Biological Bulletin 20:310-331. Della Valle, P. 1914. L’inibizione della rigenerazione del capo nelle planarie mediante la cicatrizzazione. Archivio Zoologico Italiano 7:275-311. The cause of injury probably was non-lethal predation. Linckia laevigata was reported as toxic to predators by Rideout (1975) [Toxicity of the asteroid Linckia laevigata (L.) to the damselfish Dascyllusaruanus (L.). Micronesica J. Coll. Guam 11. 1534]; on the other hand it was listed as non-toxic but protected by a very hard skeleton by Bakus (1981) [Chemical defense mechanisms on the Great Barrier Reef, Australia. Science 211:497-499]. A convincing description of toughness as defense against predation in a related starfish was given by Monks (1904). For kaleidoscopic embryology see also Dawkins (1995 A survival machine, pp. 74-95 in John Brockman, ed., The third culture: beyond the scientific revolution, Simon & Schuster, also 1996 Touchstone). Distal signaling: Midlife abrupt cessation of the ambulacral plate series in one growth zone of an echinoid supports the idea that distal signaling applies individually to each primary plate series; the interambulacral plates of the growth zone were not affected and continued up to the apical system (L.G. Zachos, "From whence cometh the plates: the ocular plate rule and Paleozoic echinoids", poster presentation at the June 2009 North American Paleontology Conference, Cincinnati, OH). Postscripts that are beyond the scope of the manuscript: Invariance of positional information along the starfish arm could be tested if it is possible to cut out a section of arm and graft the arm tip to the arm stump: my prediction is that the graft would heal without stimulating intercalary regeneration to replace the cut out portion. Likewise it could be tested if the cut out portion of arm is grafted back into place but in reverse proximo-distal orientation: my prediction is that the graft would heal without stimulating intercalary regeneration. Some additional papers on starfish regeneration: FAN Tingjun, et al., 2011, Patterns and cellular mechanisms of arm regeneration in adult starfish Asterias rollestoni Bell, Journal of Ocean University of China (Oceanic and Coastal Sea Research) 10(3):255-262. Preprint of paper by Daiki Wakita, H. Aonuma, and S. Tochinai, 2019: : Arm swapping autograft shows functional equivalency of five arms in sea stars -- doi: : Positional information in the starfish arm concluded symmetrical.

Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 2007. Isolated ossicles of the Family Eospondylidae Spencer et Wright, 1966, in the Lower Devonian of Bohemia (Czech Republic) and correction of the systematic position of eospondylid brittlestars (Echinodermata: Ophiuroidea: Oegophiurida). Acta Musei Nationalis Pragae, series B Historia Naturalis 63(1):3-18. MPRI contribution No. 1. For additional mention of Eospondylus in the Hunsrück Slate see Südkamp (1985, 1995). Further to Eospondylus on crinoids (p. 8) see Südkamp (1995); see also Kühl, Bartels, Briggs, and Rust (2012: fig. 39. Visions of a vanished world: the extraordinary fossils of the Hunsrück Slate. Yale University Press). Concerning autotomy in Eospondylus, see Kühl et al. (2012: fig. 119). Further to examples of current research (p. 6), and further to ventral and dorsal arm coiling (p. 8), Emson & Wilkie (1982. The arm-coiling response of Amphipholis squamata (delle Chiaje). In Echinoderms: Proceedings of the International Conference, Tampa Bay, ed. ed. J.M. Lawrence, 11-18, Rotterdam, A.A. Balkema) reported numerous significant observations, including the complication that the dorsal and ventral longitudinal muscles have internal and external portions that insert differently and act separately. See also the poster abstract by K. Boos, U. Nettelmann, and B. Oppermann: Effects of a chemical stimulus on the arm-coiling response in Amphipholis squamata (Delle Chiaje, 1828) (Ophiuroidea, Echinodermata), p. 90 in 14th International Echinoderm Conference, Brussels, 20-24 August 2012, Conference Booklet. Further to the postscript on an eospondylid ophiuroid of eyelet-type in the Arkona Shale, note similarities of preservation and fauna between the Arkona Shale and the Hunsrück Slate [see e.g.: Úna C. Farrell and Derek E. G. Briggs, 2007, A pyritized polychaete from the Devonian of Ontario; Proc. R. Soc. B 274:499–504. the polychaete in this paper was collected by Michael and John Topor and donated by them to UMMP]. Further to the eospondylid fossil record: Triassic Ophioflabellum (Donofrio & Mostler, 1977) lateral plates were reinterpreted as Eospondylidae by Thuy, Hagdorn, & Gale (2017 Paleozoic echinoderm hangovers: waking up in the Triassic. Geology 45:531-534, doi:10.1130/G38909.1). This identification of the figured specimens subsequently doubted and partially revised to "furcasterid protasterid" by Hunter & McNamara (Geology Nov. 2017, doi:10.1130/G39575C.1). But Thuy et al. replied and supported their original determinations as eospondylid lateral arm plates (Geology Forum November 2017; doi:10.1130/G39684Y.1)

Hotchkiss, F.H.C., and A. Glass. 2006. Bdellacoma in the Hunsrück Slate (Lower Devonian, Germany): Reidentification of Urasterella verruculosa (Asteroidea, Bdellacomidae). Poster presentation at the 12th International Echinoderms Conference. Poster [approx. 16MB PDF]. MPRI contribution No. 2. Proceedings of the conference are dated 2010 but my copy arrived 18 November 2009. Pp. 15-21 in Echinoderms: Durham, Proceedings of the 12th International Echinoderm Conference, Durham, New Hampshire, USA, 7-11 August 2006, L.G. Harris, S.A. Böttger, C.W. Walker & M.P. Lesser (eds.), CRC Press/Balkema,Leiden, 679 pp. Correction: Genus level of classification of Bdellacoma appears to date from Bronn (1860:288) rather than from Dujardin & Hupé (1862:438). Additional Bursulella fossils: Middle Devonian of western Manitoba, Canada -- see Braun & Mathison [1986. Mid-Givetian events in western Canada: The Dawson Bay -- Watt Mountain -- "Slave Point" interlude. Bulletin of Canadian Petroleum Geology 34(4):426-451, see plate 1 fig. 14; specimens lost/destroyed in a mail-handling accident (March 2012)]. Upper Ordovician erratics on Gotland Island -- see Schallreuter & Hinz-Schallreuter (2008). Also found in Ordovician microfauna from the Mishina Gora section, Russia (Tinn, Meidla, Ainsaar, Rubel & Dronov, 2011, Joannea Geol. Palaeont. 11:204; also in 8th Baltic Stratigraphical Conference Abstracts, 28 August-1 September 2011, Latvia). Additional Bd. vermiformis Leintwardine localities are Mocktree Quarry (Huxley & Etheridge 1865:74; Newton 1878:117) and Martin’s Shell Quarry (Spencer 1940:529). Stratigraphic and paleoenvironmental analysis of Leintwardine localities is in Hawkins & Hampton (1927), Whitaker (1962), Goldring & Stephenson (1972) and Gladwell (2003). The Leintwardine beds are interpreted as fine grained submarine channel deposits. Additional Bd. vermiformis materials examined: BMNH 47952, E1480 (has six mouth corners and six rays, one of which is small; figured by Blake 2013:fig. 5.3), E13958 (mentioned by Goldring & Stephenson 1972:621, "long arms drawn out"), E20242, E20243, E20245; also E52970a,b, through E52972a,b (from Martin’s Shell Quarry, but indistinct detail). USNM Springer Room: four Leintwardine specimens, USNM 92645 (qty. two), USNM 603889, and 603890. Two specimens labeled ‘Bdellacoma vermiformis’ are reidentified as Antiquaster magrumi: BMNH 40296b (see Hotchkiss 1976) and BMNH 40300. [Comment on BMNH 40296a and 40296b: numbering like this often indicates part and counterpart halves of the same fossil, but that is not the case here (RPS Jefferies, 1973 letter); 40296a is Bdellacoma, 40296b is Antiquaster.] Leintwardine specimens noted but not seen: Spencer (1940:529): Manchester University Museum no. L.11021; Royal Scottish Museum materials (see National Museums Scotland below); Geological Survey and Museum, London, nos. 54748, 54892-54894; Ludlow Museum nos. i, xii, xiii; and Ipswich Museum unspecified. Huxley & Etheridge (1865:74) and Newton (1878:117) catalogued multiple specimens in the Museum of Practical Geology [forerunner of the Geological Survey Museum], but none are indicated as type specimens by Allen (1902). Salter (1873:163) catalogued a specimen in the Geological Museum of the University of Cambridge. National Museums Scotland has multiple specimens: NMS G.1882.65.152.1 through NMS G.1882.65.152.6 (formerly Royal Scottish Museum RSM numbers; NMS G.1882.65.152.3 is the neotype selected by Spencer) (Lyall Anderson personal communication December 2003). “Wenlock Herefordshire Lagerstätte” materials of Sutton et al. (2005): Nodules with calcite infills that preserve high-fidelity details of soft tissues, but stereom has recrystallized and plate boundaries are not apparent. University Museum of Natural History, Oxford, OUM C.29572-C.29585. Specimens OUM C.29572-C.29573 serially ground and computer reconstructed with “virtual dissection” image capability. Family classification of Bdellacoma: Spencer (1951:111) as Family Phragmactidae (Ophiuroidea); Spencer & Wright (1966:U81) as Family Bdellacomidae (Ophiuroidea). Urasterella verruculosa was described by Lehmann (1957:131-134) and has been mentioned/recognized by Spencer and Wright (1966: p. U71 and Fig. 64:6e), Kutscher (1967, 1970), Bartels and Brassel (1990:142-144) who gave a concise description and illustrated a specimen with only four arms (fig. 127a), and Bartels, Briggs and Brassel (1998) who figured a five-rayed specimen (Fig. 183) and mentioned the four-rayed occurrence (p. 227). Arms taper in a 2 cm specimen figured by Rudolf Stanzel (2014. Bundenbach -- Teil 2 -- Jugendformen. Der Steinkern ISSN-1867-8858, Heft 16:14-21, see fig. 17). The madreporite has now been seen in Bd. vermiformis by Gladwell (2005 thesis; also Gladwell in Lewis et al. 2007). About ambulacral versus adambulacral plates: Lehmann thought that the ambulacrals of Bd. verruculosa are armed with delicate spines, and specimen YPM 217481 gives this appearance. Likewise Sutton et al. (2005:1004) saw spine-bearing transverse ridges associated with the ambulacral groove. We presume that ambulacral plates do not carry spines, and interpret that the spine-bearing adambulacral plate is ventrally superposed on the ambulacral plate: i.e. superposed on the transverse bar of the T-shaped ridge. YPM 217481 then indicates that the superposed adambulacral plates could slide inward to better cover the ambulacral groove with spines. About pedicellariae: The pedicellariae of Bdellacoma are remarkable and deserve further study. Papers relevant to such a study could include: Breton, Gérard. 1996. Les pédicellarires alvéolaires sont-ils arrivés chez les Goniasteridae au Crétacé par voie épidémique? Bull. Soc. zool. Fr. 121(1):87-92. Pedicellariae described also in: Kesling, Robert V. 1982. Arkonaster, a new multiradiate starfish from the Middle Devonian Arkona Shale of Ontario. Contrib. Mus. Paleont. Univ. Mich. 26(6):83-115. On p. 144 of Echinoderm Research vol. 1, Andrew Campbell, Form and function of pedicellariae, note to Table 2 says: “Some starfish which as adults lack pedicellariae bear them on larvae; i.e. some astropectinids (Downey, personal communication)”. See also: Turner, R.L., J.M. Boucher, and M.L. Wittenrich. 2008. A new kind of pedicellaria of echinoderms [abstract]. Fifth North American Echinoderm Conference, Florida Institute of Technology, Melbourne, Florida program with abstracts. And watch for a paper on Helianthaster by D.B. Blake in 2009. Addendum to literature: Lamont (1947, Gala-Tarannon Beds in the Pentland Hills, near Edinburgh, Geol. Mag. 84(4):193-208, on p. 203, Bdellacoma vermiformis in the Pentland Hills, based on Spencer 1940);Thayer (1972 thesis; 1974); Hotchkiss & Glass (2008 in 4th Workshop of German & Austrian Echinoderm Research book of Abstracts).

Hotchkiss, F.H.C. and R. Haude. 2004. Observations on Aganaster gregarius and Stephanoura belgica (Ophiuroidea: Ophiolepididae) (Early Carboniferous and Late Devonian age). Pp. 425-431 In Heinzeller & Nebelsick (eds.), Echinoderms:München. Taylor & Francis Group, London. The purpose of the research was to elucidate the structure of the oral frame, the structure of the distal parts of the rays, and the status of radial shields. It was noted that this was partly accomplished with new material. More specimens need to be examined to substantiate and expand the findings, especially any observations on mouth papillae. MPRI 0077 is a slab that has five fairly intact disks and many isolated portions of arms of A. gregarius. The slab is now in the Yale Peabody Museum as YPM IP 521654 - 521658. YPM IP 521654 preserves one arm out to the 18th arm segment in under-surface view. This arm confirms that the under arm plates of Aganaster abruptly cease after the 15th arm segment, and that podial pores perforate the lateral arm plates beginning on the 16th arm segment. This arm increases the number of observations of this question to N = 3. YPM 38474 is an arm fragment that is 14 segments long and shows the abrupt cessation of under arm plates, but the arm segment numbers for this fragment are not determinable. Addendum to observations on A. gregarius: The oral plates do not have lateral wings (Hotchkiss & Haude 2004 figures 1D, 1G). An error on pages 429-430: Archaeophiomusium is corrected spelling. Open questions: Does A. gregarius have disc granules? To be noted in passing: Matsumoto (1913, 1929) classified A. gregarius in his new Order Liparophiurida (with additional content), but this has not been adopted/supported by subsequent research. Pearse (1947. Zoological names - a list of phyla, classes, and orders) listed Order Aganasterida H.L. Clark 1939. I have not found the 1939 H.L. Clark paper, in spite of earnest searching. The Order Aganasterida is listed also by Boettger (1952) and by Blackwelder (1963). Additional specimens of A. gregarius: Eight slabs donated November 2004 to the Natuurhistorisch Museum Maastricht, registration numbers NHMM 2004 156 through NHMM 2004 163. Seven slabs donated October 2010 to the Geoscientific Museum, George August University of Goettingen (MPRI 0250 through 0256). New literature: Thuy, B., Kutscher, M., and Płachno, B.J. 2015. A new brittle star from the early Carboniferous of Poland and its implications on Paleozoic modern-type ophiuroid systematics. Acta Palaeontologica Polonica 60 (4): 923–929. New literature: O’Hara, T. D., A. F. Hugall, B. Thuy, and A. Moussalli. 2014. Phylogenomic resolution of the Class Ophiuroidea unlocks a global microfossil record. Current Biology 24, 1874–1879. New literature: Ben Thuy and Sabine Stöhr. 2011. Lateral arm plate morphology in brittle stars (Echinodermata: Ophiuroidea): new perspectives for ophiuroid micropalaeontology and classification. Zootaxa 3013: 1–47.

Hotchkiss, F.H.C. 2000. On the number of rays in starfish. American Zoologist 40:340-354. Correction to the statement concerning holothurians on p. 340: the tentacles of holothurians correspond to the rays of other echinoderms, and holothurians have FIVE-PLUS organization of rays/tentacles around the mouth (Hotchkiss 1998:rays as appendages, p.208). To p. 342 on Paleozoic aberrant ray number add two of over 400 Devonaster eucharis found 4-rayed (Clarke 1912, Bull. NYSM No. 158, pp. 44-45 + pls). Further to "Culcita tetragona" on p. 342: appears also in Sedgwick's (1908, p. 167) vol. 3 of "A student's textbook of zoology". Further to 6-rayed Aleutiaster schefferi on p. 342: reclassified from Ganeriidae to Solasteridae by Gale (personal communication April 2017; see also Jewett et al., Sea stars of the nearshore Aleutian Archipeligo, pp. 144-172, in Stelleer & Kerr, eds., Diving for Science 2012, Proc. Am. Acad. Underwater Sciences 31st Symposium, Dauphin Island, AL). With this revision the Ganeriidae are strictly 5-rayed. The 8-rayed specimen of Luidia senegalensis on p. 345 is deposited in the Smithsonian National Museum of Natural Histsory as USNM 1112625. In 2004 Balser reported that L. senegalensis metamorphosed with 9 rays present, but did not comment on the work of Komatsu et al (1991) that reared L. semegalensis to metamorphosis and observed only five rays at metamorphosis [Balser, E.J. 2004. pp. 3-9 in Echinoderms-Munchen, 11th IEC Proceedings]. Further evidence that tetramery at metamorphosis is the result of teratological incomplete development (p. 349) comes from an Arbacia punctulata that developed 4-part symmetry while its genetically identical twin developed 5-part symmetry (the parents had typical 5-part symmetry; Marcus, N.H. 1979. Developmental aberrations associated with twinning in laboratory-reared sea urchins. Developmental Biology 70:274-277). Likewise in the ophiuroid Amphipholis squamata, deviations from pentamerism are not heritable but are a consequence of environmental perturbations on the metamorphosis of larvae and/or abnormal regeneration of arms [Dupont, S. & J. Mallefet. 2002. Abnormal forms in the brittle-star Amphipholis squamata: a field study. J. Mar. Biol. Ass. U.K. 82:491-493]. Exposure of Heliocidaris erythrogramma to NiCl2 perturbed development; outcomes included symmetry variants from biradial to decaradial (33% nonpentaradial) [Minsuk, S.B., F.R. Turner, M.E. Andrews & R.A. Raff. 2009. Axial patterning of the pentaradial echinoderm body plan. Dev Genes Evol 219:89-101]. Some additional references: 2000: Sanchez, P. The sequence of origin of the postmetamorphic rays in Heliaster and Labidiaster (Echinodermata: Asteroidea). Revista Chilena de Historia Natural 73:573-578. 2001: Sumida, P.Y.G., P.A. Tyler & D.S.M. Billett. Early juvenile development of deep-sea asteroids of the NE Atlanntic Ocean, with notes on juvenile bathymetric distributions. Acta Zoologica (Stockholm) 82:11-40 [Brisingella coronata with disk sizes from 2.08 mm to 5.38 mm, the number of arms varied between 8 and 10, no relation of arm number to size was noted]. 2004: Balser, E.J. And then there were more: cloning by larvae of echinoderms. pp. 3-6 in Echinoderms: Munchen, Proceedings of the 11th IEC [reported that Luidia senegalensis metamorphoses with 9 rays present]. 2007: Herringshaw, L.G., M.P. Smith and A.T. Thomas. Evolutionary and ecological significance of Lepidaster grayi, the earliest multiradiate starfish. Zoological Journal of the Linnean Society 150:743-754. 2008: Bos, A.R., J.C.E. Alipoyo, L.T. Cardona, G.S. Gumanao, and F.N. Salac. Population structure of common Indo-Pacific sea stars in the Davao Gulf, Philippines. UPV J. Nat. Sci. 13 Suppl. pp. 11-24 [irregular number of rays may be strongly affected by predation]. 2009: Villier, L., S. Charbonnier & B. Riou. Sea stars from Middle Jurassic Lagerstätte of La Voulte-sur-Rhône (Ardčche, France). Journal of Paleontology 83(3):389-398 [describes Decacuminaster solaris n.sp. with 17 and 18 arms, Order Velatida, cf. Myxasteridae]. 2010: Lawrence, J.M., and W. Avery. Bilateral symmetry of the rays of small Luidia senegalensis (Echinodermata: Asteroidea). Florida Scientist 73(1):27-30 [A small specimen with five rays of R = 7.1mm has four rays of R = 5.1mm, one each between the larger rays; there is no sign of injury, indicating that this is the method of ray addition in the post-metamorphic 5-rayed L. senegalensis imago.] 2010: Shibata, D., Y. Moriyama, and M. Komatsu. Development of the fissiparous and multiarmed seastar, Coscinasterias acutispina (Stimpson). pp. 479-486 In "Echinoderms: Durham" (eds. L.G. Harris, S.A. Bottger, C.W. Walker and M.P. Lesser) Taylor & Francis Group, London [There is a pause between the brachiolaria with 5 hydrolobes and the brachiolaria with 6 lobes; juvenile has 6 rays and one madreporite at metamorphosis; schizogony into two individuals each with 3 arms; new arms come in as 2+2+1; total after first division is 8 arms of adult]

Hotchkiss, F.H.C. 2000. Inferring the developmental basis of the sea star abnormality “double ambulacral groove” (Echinodermata: Asteroidea). Revista Chilena de Historia Natural 73:579-583. LINK TO PAPER The specimen (p. 582) from Marblehead, MA of Asterias vulgaris with double ambulacral groove on one of only four rays is deposited in the collections of the California Academy of Sciences, CASIZ 173969. The specimen (p. 581) from a student at Lehigh University of Asterias forbesi with double ambulacral groove on one of its five rays is deposited in the US National Museum, Smithsonian Institution, echinoderm collection, USNM 1082952. The Zoological Record for 1904, p. 34, mentions fusion of rays in Ctenodiscus, referencing Michaailovskij, M. 1904. Die Echinodermen der zoologischen Ausbeute des Eisbrechers “Jermaks” von Sommer 1901. Annuaire Mus. St. Petersb. ix, pp. 157-188. The research collection of sea stars of H. Barraclough Fell contained a Pentagonaster pulchellus with double ambulacral groove. This collection is now at the US National Museum. Living specimen of Pisaster with double ambulacral groove, found May 2017, by Prof. Eric Sanford, Bodega Marine Laboratory, CA; specimen deposited in the collection of the California Academy of Sciences under CASIZ-224115.

Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 1999. Isolated skeletal ossicles of a new brittlestar of the Family Cheiropterasteridae Spencer, 1934 (Echinodermata: Ophiuroidea) in the Lower Devonian of Bohemia (Czech Republic). Journal of the Czech Geological Society 44:189-193. To the occurrences of the family Cheiropterasteridae, add the Silurian and Lower Devonian of Bolivia (Branisa, Leonardo. 1965. Los fosiles guis de Bolivia I. Paleozoico. Index Fossils of Bolivia I. Paleozoic. Servicio Geologico de Bolivia, Boletin No. 6, 282 pp., 80 pls. [?Loriolaster sp. p. 108 pl. xxii fig. 1] [?L. cf. L. mirabilis p. 128 pl. xxxii fig. 9]). Also Gladwell [in Lewis et al. 2007 A field guide to the Silurian Echinodermata of the British Isles: Part 1 Eleutherozoa and Rhombifera. Scripta Geologica 134:27-59, on page 35 and plate 1 fig. 7] reported Loriolaster sp. nov. from Leintwardine (see Hotchkiss 1978 p. 539 where this was previously indicated). The replacement name for Hexura Spencer, 1950, is Hexuraster Jell & Theron, 1999. Milwaukee Public Musem specimen no. 25640 of Loriolaster mirabilis, in oral view, displays a definite, low-profile, button-like madreporite with meandering madreporiform markings (madreporite not previously seen so clearly).

Hotchkiss, F.H.C., Rudolf J. Prokop and Václav Petr. 1999. Isolated vertebrae of brittlestars of the Family Klasmuridae Spencer, 1925 (Echinodermata: Ophiuroidea) in the Devonian of Bohemia (Czech Republic). Journal of the Czech Geological Society 44:329-333. The Narodni Museum specimen number for Antiquaster sp. material from the Renault Formation at Waterloo, Illinois, is S 04784. Further to the occurrences of the family Klasmuridae, add isolated ossicles in the Devonian of Poland by Bockzarowski (2001, especially pp. 13-20) with descriptions of new species of Pectenura: P. excubitor, P. formosa, P. pecten, and P. senta. Also isolated lateral element from a borehole, Devonian of Germany, by Haude (2004, on p. 244) [Haude, R. 2004. Cour. Forsch.-Inst. Senckenberg 251:237-251]. Regarding Antiquaster magrumi add: Kesling & Chilman (1975, see pp. 168, 176, and plate 39) [UMMP Papers on Paleontology No. 8]. For redescription of Protasteracanthion and P. primus, and Antiquaster made a synonym of Protasteracanthion, see Alexander Glass (2006. Ph.D. Thesis. The brittle star fauna of the Hunsruck Slate and a phylogeny of the Paleozoic Ophiuroidea. Univ. Illinois at Urbana-Champaign. For abstract see Dissertation Abstracts International 67(11B):6270].

Hotchkiss, F.H.C. 1998. Discussion on pentamerism: The five-part pattern of Stromatocystites, Asterozoa, and Echinozoa. Pp. 37-42 In R. Mooi and M. Telford (eds.) Echinoderms: San Francisco. A.A. Balkema, Rotterdam. Postscript notes: Nichols (1967b) noted that the crystallographic alignment hypothesis of Cockbain (1966) is not consistent with c-axis orientations in echinoid genital plates; clarified and expanded arguments that support the suture-line hypothesis. There is much additional literature relevant to this topic that could not be fitted into the original paper; also there is literature that has come out since the paper. 1889. Bather, F.A. Trigonocrinus, a new genus of Crinoidea, from the "Weisser Jura" of Bavaria; with the description of a new species, T. liratus.--Appendix. Sudden deviations from normal symmetry in Neocrinoidea. Quarterly Journal of the Geological Society for February 1889, pp. 149-171, pl. 6 [p. 166 pentamery persists through natural selection based on mechanical principles; in pentamerous calyx every line of weakness is met halfway by a solid plate]. 1915. Clark, A.H. A monograph of the existing crinoids. The comatulids. USNM Bull. 82, vol. 1, part 1, 406 pages [p. 148: sutures are lines of weakness; with an odd number of radial divisions the sutures do not line up and the construction is stronger; the relation between the bilateral crustacean type and the pentaradiate echinoderm type is explained and provides explanation for the establishment of pentaradial symmetry]. 1921. Clark, A.H. The echinoderms as aberrant arthropods. Smithsonian Miscellaneous Collections 72(11):1-20 [p. 7: the echinoderm body is composed of five half metameres; the anterior and posterior ends of the series of half metameres join; pp. 18-20: lengthy quote from book by William Patten, 1912, "The evolution of the vertebrates and their kin", who proposed the same idea, see pages 421-430; the quote refers to a marked asymmetry as a frequent abnormality in Xiphosura, the context suggesting half metameres; an abnormality resembling this concept is described by T. Itow, R.E. Loveland & M.L. Botton, 1998, Developmental abnormalities in horseshoe crab embryos caused by exposure to heavy metals, Arch. Environ. Contam. Toxicol. 35:33-40, Fig. 25 "half embryo"]. 1928. Bather, F.A. The fossil and its environment. Quarterly Journal of the Geological Society of London 84(2):lxi-xcviii [pp. lxxi-lxxii present his ideas on the origin in Pelmatozoa of a primitive trisactiny followed by a forking of the lateral rays; a pentactiny superimposed upon a trisactiny; states the advantage of tripartite and quinquepartite over quadripartite: each line of weakness abuts on a mass of strength; calling it the principle of bonding]. 1930. Clark, A.H. The new evolution: Zoogenesis. The Williams & Wilkins Co., Baltimore, xiv+297 pp. [Fig. D on p. 250 on animal symmetries includes "the pseudoradial symmetry of an echinoderm -- really bilateral symmetry with the axis curved in a circle, half of the five segments of the body failing to develop in the adult"]. 1975. Strathman, R.B. Limitations on diversity of forms: branching of ambulacral systems of echinooderms. The American Naturalist 109:177-190. 1976. Frest, T.J., and H.L. Strimple. Evolutionary and paleoecologic significance of abnormal Platycystites cristatus Bassler (Echinodermata: Paracrinoidea). J. Wash. Acad. Sci. 66(4):221-228 [1-1 is primitive; 1-1-1 and 2-1 and 1-2 and 2-1-2 are derived]. 1987. Anderson, D.T. Developmental pathways and evolutionary rates. In: Rates of Evolution, K.S.W. Campbell & M.F. Day (eds.), Allen & Unwin, Sydney [p. 146 pentamerous symmetry determined during development by a counting process]. 1993. Mae-Wan Ho and Peter T. Saunders. Rational taxonomy and the natural system with particular reference to segmentation. Acta Biotheoretica 41:289-304 [on p. 294: they interpret 3, 4, and 6-fold aberrant echinoids and asteroids of Raff & Kaufman (1983) as a transformation set from an underlying dynamical structure (= low mode multifurcation)]. 1993. Trinajstić, Nenad. The magic of the number five. Croatica Chemica Acta 66(1):227-254 [p. 236 pentagonal structure in echinoderms suited to sedentary habits and to facing threats from all sides; plus a broad discussion of the number five in many contexts]. 1998. Tolić, Iva Marija and Nenad Trinajstić. The number five in biology. Periodicum Biologorum 100(3):259-266 [Fivefold symmetry in animals, plants, viruses, biomolecules, crystals (quasicrystals)]. 1998. Prokop, R. and V. Petr. How the Echinoderm Changed Its Symmetry, or On the search for common mechanisms to generate the archetypal forms. Bulletin of the Czech Geological Survey, vol. 73, n. 4, 351-354. 1999. Torres Hernanz, M., Gil Cid, M.D., and Dominguez Alonso, P. Stable multimodal fivefold patterns: the case of Rhodocrinitids. Coloquios de Paleontologia 50:117-25 [in Spanish; mathematical approach]. 2001. Changizi, M.A. The economy of the shape of limbed animals. Biological Cybernetics 84:23-29 [wire-minimization principles, animal limb number, and body-to-limb proportion; predicts six or more rays for Asteroidea]. 2002. Barrio, R.A., P.K. Maini, J.L.Aragón and M. Torres. Size-dependent symmetry breaking in models for morphogenesis. Elsevier Physica D 168-169(2002):61-72 [discusses de novo appearance and control of pentagonal symmetry of echinoids]. 2002. Torres, M., Aragon, J.L., Dominguez, P., and Gil, D. Regularity in irregular echinoids. J. Math. Biol. doi 10.1007/s002850100126 Springer-Verlag [eutactic star morphospace]. 2006. Parsley, R.D. and Y. Zhao. Long stalked eocrinoids in the basal Middle Cambrian Kaili Biota, Taijiang County, Guizhou Province, China. J. Pallaeont. 80(6):1058-1071 [pp. 1069-1070 discuss the ontogeny and phylogeny of ambulacral symmetry, from a 1-1 pattern to a 2-2 pattern to a 2-1-2 pattern]. 2006. Simon Conway Morris. Darwin's dilemma: the realities of the Cambrian 'explosion'. Philosophical Transactions of the Royal Society B, 361:1069-1083 [p. 1075 prepentameral sequence of vetulocystids, stylophoran cornutes, cinctans and solutes; subsequent pentameral symmetry may reflect the transition to sessility]. 2010. López-Sauceda, J., and J.L. Aragón. Pentamerism and modularity in sea urchins. Tip Revista Especializada en Ciencias Quimico-Biológicas 13(2):121-125 [modular analysis provides clues to advantages of pentamrism]. 2012. Liang Wu et al. The advantages of the pentameral symmetry of the starfish []. 2012. Parsley, R.L. Ontogeny, functional morphology, and comparative morphology of lower (Stage 4) and basal middle (Stage 5) Cambrian gogiids, Guizhou Province, China. Journal of Paleontology 86(4):569-583 [p. 571 probably an initial 1-0-1 ambulacral pattern; followed by a 2-0-2 pattern; later a single ambulacral branch forms at the front of the mouth to produce a 2-1-2 pattern; the presage of pentamery of other echinoderm groups]. 2012. G.D. Webster and S.K. Donovan. Before the extinction – Permian platyceratid gastropods attached to platycrinitid crinoids and an abnormal four-rayed Platycrinites s.s. wachsmuthi (Wanner) from West Timor . Palaeoworld Volume 21, Issues 3–4, December 2012, Pages 153–159 [on p. 154 they use the term quadrilateral symmetry and offer the possibility that it is a genetic defect; FHCH cannot agree because this claims that ray number has heritable variation and requires that pentamery be maintained by natural selection based on superior fitness; the observed extreme fidelity to pentamery is better explained as a developmental constraint].

Hotchkiss, F.H.C. 1998. A "rays-as-appendages" model for the origin of pentamerism in echinoderms. Paleobiology 24:200-214. A gentle criticism of the idea that there are analogies between appendages of bilateral forms and the rays of echinoderms is thoughtfully presented by R. Mooi, B. David & G. Wray, 2005, Arrays in rays: terminal addition in echinoderms and its correlation with gene expression, Evolution and Development 7(6):542-555. I accept their comments and claim only the analogy; the phrase rays-as-appendages seemed apt because Baterson's rule of duplication was discovered from analysis of duplicated appendages, and does not apply to axial structures; however Bateson's rule is not limited to appendages (it operates also in ciliate protozoa). Further to the mention on p. 203 of the paper by Hinegardner (1975): The report of loss of symmetry control and the rearing of four-part "square" Lytechinus pictus challenges the working hypothesis of locked-in pentamerism. Therefore it is important to mention that these are highly inbred lines with high mortality as larvae and at metamorphosis, facts which suggest that the square imagos are the result of teratological incomplete development. I do not agree with D.G. Stephenson's (Proc. European Colloq. Echinoderms,Brussels, 1979) interpretation that Hinegardner's square sea urchins are the result of loss of symmetry as soon as domestication relaxes the constraints of natural selection; nor Stephenson's corollary that pentamerism must be extrtemely adaptive for echinoids. Stephenson also presented the idea that "pentamerism is an ancestral character retained because any disadvantages of pentamerism are less than the disadvantages of changing it", and claimed that "this possibility has been ruled out by Hinegardner's (1975) breeding experiments." I propose instead that the loss of symmetry reported by Hinegardner be interpreted along the lines described by John M. Opitz (1985. The developmental field concept. American Journal of Medical Genetics 21:1-11, see Final generalizations on pp. 9-10): "all primary malformations qua developmental field defects are causally nonspecific"; "most will be shown to be causally heterogeneous"; "the causes are many, but the final common developmental pathways are few"; "most primary malformations are anomalies of incomplete formation"; "most field defects are multifactorially determined (not 'inherited'), hence have a low empiric recurrence risk"; "because oligogenes of major effect do play a role in the developmental history of all fields, sooner or later Mendelian inheritance of a primary malformation should be expected and observed"; "most field defects occur per se but can be and frequently are components of complex syndromes, including aneuploidy syndromes"; "the epigenetically hierarchichical nature of field development is responsible for the fact that, regardless of how small or limited the embryonic primordium, so long as it is still a field, its development can be altered genetically and environmentally." A further reference to diatoms with pentagonal symmetry: P.A. Simms & R. Ross. 1990. Triceratium pulvinal and T. unguicularum, two confused species. Diatom Research 5(1):155-169 [figs. 13-16 of T. unguiculatum forma quinquangulum]. Further to favoring ray identifications based on position of the place of hydrocoel closure (p. 201): add both Sedgwick and MacBride (Sedgwick, 1909, vol. 3, Student’s textbook of zoology, p. 119 footnote, citing MacBride). Further to the matrix of ray homologies and especially concerning the entry for Holothurians, and not in my table [because I did not see this Semon (1888) reference when I wrote my paper], the relation of pentameral symmetry of the adult to the bilateral symmetry of the larva was tracked in diagrams by Semon (1888: plate 1A) concerning the development of Synapta digitata. His diagrams show that the larval anus (P = periproct) is carried through to the adult and lies ventrally in the plane of hydrocoele closure (R = place of ring closure). This places both R and P in interradius IV/V [Loven numerals assigned from place of R in echinoid development]. In subsequent Synapta digitata development the anus moves to the distal end of the holothurian where assignment to an interradial position is not evident. [Semon, R., 1888, Die Entwickelung der Synapta digitata und die Stammesgeschichte der Echinodermen, Jenaischen Zeitschrift fur Naturwissenschaft 22(N.F. XV): 135 pp., 7 plates.] Further to the matrix of ray homologies and especially concerning the entry for Luidia: My primary source on the location of hydrocoel closure and the madreporite in Luidia is Bury 1895 Fig. 21 and text pp. 67-68 [Bury 1895 The metamorphosis of echinoderms. Q. J. Microsc. Sci. n.s. 38:45-136 + pls. 3-9] [this source is mentioned in the caption to my matrix of ray homologies, but change "Fig. 22" to "Fig. 21"]. The observations of Bury were latched onto by F.A. Bather [1915 Studies in Edrioasteroidea IX. The genetic relations to other echinoderms. Geol. Mag., n.s., dec. VI, vol. 2, pp. 393-403]. On page 401 “It is important to notice here that in an Asteroid larva (Bipinnaria asterigera) described by Bury, the [hydrocoel] closure does still take place in the M plane” and mentioned further on p. 402. Mentioned again by Horstadius [1939 Uber die Entwicklung von Astropecten aranciacus L. Pubb. Staz. Zool. Napoli 17(2):221-312]. On p. 276 he says that in all known starfish (including Astropecten) the madreporite is between hydrocoel lobes I-II, except in Luidia it is between lobes V and I. On p. 278 he appears to give his own confirmatory observations on this point in recounting his work on Luidia ciliaris. NOTE NOTE NOTE – his system of Roman numerals is customary numbering of the linear series of hydrocoel lobes as seen in developing imago of metamorphosing larva, so is different than in my matrix. His Roman I is my edrio and asteroid D; his Roman V is my edrio and asteroid C; his Roman II is my edrio and asteroid E. Further again to data on Luidia: Bilateral symmetry of lengths of rays has been definitively determined, with the madreporite and ring-closure on this axis. See Lawrence, Pomoroy & Trowbridge [2011. Symmetry of the rays of Luidia clathrata (Echinodermata: Asteroidea). abstract from 6th North American Echinoderm Conference, 14-19 August 2011, Rosario Beach Marine Laboratory, Anacortes, WA, Gulf of Mexico Science 29(2):147]. Further again to the matrix of ray homologies, and for fairness of presentation, let me mention that there is not yet a concensus on ray homologies: for a contrasting opinion on ray homologies see V.B. Morris et. al. (2009, Development of the five primary podia from the coeloms of a sea star larva: homology with the echinoid echinoderms and other deuterostomes. Proc. R. Soc. 276:1277-1284; see especially p. 1283). Further to echinoids in the matrix of ray homologies: hydrocoele ring closure between ambulacra IV and V was confirmed in Peronella japonica by Tsuchimoto et al. (2011:2436) [Tsuchimoto, J., T. Yamada, and M. Yamaguchi. 2011. Unusual coelom formation in the direct-type developing sand dollar Peronella japonica. Developmental Dynamics 240:2432-2439.] Further to p. 204 morphological principles see Hermann Weyle book "Symmetry" pp. 27-28 (1952, reprinted 1989 Princeton Science Library) on origination and retention of bilateral symmetry, also on asymmetry and helical progression (FH: think of helicoplacoids; think of sideways crabwalk; and see pp. 207--208 on the reorientation of von Ubisch's L/R into an altered A/P 2-3 pattern in irregular echinoids). Some newer papers on the development of the echinoderm body plan: 1989: Hahn, W. Symmetrie als Entwicklungsprinzip in Natur und Kunst. Koenigstein (Langewiesche). 1998: Gehring, W.J. Master control genes in development and evolution: the homeobox story. Yale University Press 236 pp [pp. 24-25 the possibility that six-armed starfish carry a homeotic mutation]. 1999: Kerr, A.M., J. Kim. Bi-penta-bi-radial symmetry: A review of evolutionary and developmental trends in holothuroidea (Echinodermata). Journal of Experimental Zoology (Mol. Dev. Ecol.) 285:93-103. 2002: Aragon, J.L., M. Torres, D. Gil, R.A. Barrio, P.K. Maini. Turing patterns with pentagonal symmetry. Physical Review E, vol. 65, 051913:1-9 [may offer a possible mechanism for inducing five-fold symmetry observed in early development of sea urchins, crinoids]; Barrio, R.A., P.K. Maini, J.L. Aragon, M. Torres. Size-dependent symmetry-breaking in models for morphogens. Physica D 168-169:61-72; Gudo, M. and M. Grasshoff. The origin and early evolution of chordates: the "Hydroskelett-Theorie" and new insights toward a metameric ancestor. Senkenbergiana lethaea 82(1):325-346. 2003: Minelli, A. The origin and evolution of appendages. Int. J. Dev. Biol. 47:573-581 [vertebrate tail and echinoderm radii -- body axis versus appendage]; Minelli, A. The development of animal form -- ontogeny, morphology, and evolution. Cambridge Univ. Press, 323 pp [p. 129 multiradiate starfish; p. 156]. 2006: Hibino, T., A. Nishino and S. Amemiya. Phylogenetic correspondance of the body axes in bilaterians is revealed by the right-sided expression of Pitx genes in echinoderm larvae, Develop. Growth Differ. 48:587-595; Swalla, B.J. Building divergent body plans with similar genetic pathways, Heredity 97:235-243. 2008: Minsuk, S.B., F.R. Turner, M.E. Andrews and R.A. Raff. Axial patterning of the pentaradial adult echinoderm body plan. Dev Genes Evol 219:89-101. 2009: Guensburg, T.E. and J. Sprinkle. Solving the mystery of crinoid ancestry: new fossil evidence of arm origin and development, Journal of Paleontology 83(3):350-364. 2011: Tsuchimoto, J., T. Yamada, and M. Yamaguchi. Unusual coelom formation in the direct-type developing sand dollar Peronella japonica. Developmental Dynamics 240:2432-2439.

Hotchkiss, F.H.C., D. M. Rudkin, and S. Anderson. 1997. A case for rearmament -- the oldest known evidence of regeneration in sea stars (Abstract). 7th Canadian Paleontology Conference, University of Saskatchewan, Saskatoon, September 26-30, 1997. [Also: Rudkin, D. M., F. H. C. Hotchkiss and S. Anderson. A case for rearmament -- the oldest known evidence of regeneration in sea stars. Royal Ontario Museum Nineteenth Annual Research Colloquium, 20 November 1997, Toronto, Abstracts of Papers, p. 1 (abbreviated abstract).] We reported a specimen of Promopalaeaster wilsonae (Raymond 1912) with two regenerating arm tips, collected by S. Anderson from the Upper Member of the Bobcaygeon Formation (Middle Ordovician: Kirkfieldian) of south-central Ontario, and now housed in the Royal Ontario Museum (ROM 53320). The Anderson fossil is the oldest [earliest] example of an asteroid with a regenerating ray, supplanting a regenerating specimen of Cnemidactis girvanensis from the Upper Ordovician of Girvan, Scotland reported by Spencer (1918 pt. III, p. 161, pl. 12 fig. 5). Arm stumps that had healed without regenerating the lost arm were described in the Ordovician starfish Urasterella ulrichi by Schuchert (1915:37).

Hotchkiss, F.H.C. 1995. Lovén's law and adult ray homologies in echinoids, ophiuroids, edrioasteroids and an ophiocistioid (Echinodermata: Eleutherozoa). Proceedings of the Biological Society of Washington [DC], vol. 108, no. 3, pp. 401 435. Lovén’s Rule in the echinoid order Cidaroida: Lovén’s Rule is evident in the pattern of alternation of the ambulacral plates that begins at the edge of the peristome in a 7 mm diameter specimen of Histocidaris elegans (A. Agassiz) figured by Mortensen (1927: Fig. 6). Reference: Mortensen, Th. 1927. Report on the Echinoidea collected by the United States Fisheries Steamer “Albatross” during the Philippine Expedition, 1907-1910. Part 1 The Cidaridae. USNM Bulletin 100, vol. 6, part 4, pp. 243-312, pls. 48-80. Lovén’s Rule in the echinoid order Temnopleuroida: in Temnopleurus apodus only the buccal plates Ib, IIb, IIIa, IVb, and Va carry tubefeet, the other buccal plates being quite rudimentary (Mrtsn 1943 Monograph III.2 p. 101); in Paratrema doederleini with only five buccal plates, the five buccal plates that are present are the plates Ib, IIb, IIIa, IVb, Va (R. T. Jackson, 1927, p. 451, fig. 8; confirmed by Mrtsn 1943 Monograph III.2 p. 274; a specimen with only four buccal plates has the A. II buccal plates missing); however in Prionechinus saggitiger with only five buccal tube feet the pattern is bbbba (Agassiz, 1881, Challanger Report, pl. VIa, fig. 12; numbering of ambulacra not stated). Lovén’s Rule in the echinoid order Echinoida: In Strongylocentrotus droebachiensis and in Psammechinus miliaris the buccal tube-feet first appear in the plates Ib, IIb, IIIa, IVb, Va (Jackson 1927, pp. 451-452; David & Mooi, 1995, The ontogenetic basis of Lovén’s Rule clarifies homologies of the echinoid peristome, pp. 155-164, in Echinoderm Research 1995, Emson, Smith & Campbell (eds), Balkema, Rotterdam; key reference on Lovén’s Rule in echinoids and other Echinodermata). Lovén’s Rule in the echinoid order Holasteroida: the largest ambulacral basicoronals in pourtalesiids definitely exhibit the aabab pattern early in ontogeny (David & Mooi 1995). Lovén’s Rule in the echinoid order Clypeasteroida: see Bruno C. Vellutini and Alvaro E. Migotto, "Embryonic, larval, and juvenile development of the sea biscuit" Clypeaster subdepressus (Echinodermata: Clypeasteroida), PLoS ONE March 2010, vol. 5, issue 3, e9654 []. Specimen USNM 376690 of "Stromatocystites walcotti" (the specimen that conforms with Lovén’s rule) is judged closely related to Kailidiscus chinensis by Zhao et. al. 2010 [Yuanlong Zhao, Colin D. Sumrall, Ronald L. Parsley and Jin Peng, Kailidiscus, a new plesiomorphic edrioasteroid from basal Middle Cambrian Kaili Biota of Guizhou Province, China. Journal of Paleontology 84(4):668-680]. Concerning glyptocystitoid rhombiferans and Lovén’s rule: according to Sumrall (2008), both glyptocystitoids and hemicosmitoids follow Lovén’s rule in the placement of brachioles [Sumrall, C. D. 2008. The origin of Lovén’s rule in glyptocystitoid rhombiferans and its beraring on the plate homology and heterochronic evolution of the hemicosmitoid peristomial border. chapter 11 (pp. 228-241) in W. I. Ausich & G.D. Webster, eds., Echinoderm Paleobiology, Indiana University Press, 456pp.]. Sumrall & Wray (2007:152-153 and fig. 4B) recognized that "in many groups of echinoderms the A, C, and E ambulacra have identical morphological characteristics, while the B and D ambulacra share a mirror-image of those characteristics, a pattern known as Lovén’s Law", but their diagram of Lovén’s Law in glyptocystitoid rhombiferans does not conform with the mirror-image portion of this definition of Lovén’s Law [Sumrall, C. D. and G. A. Wray. 2007. Ontogeny in the fossil record: diversification of body plans and the evolution of "aberrant" symmetry in Paleozoic echinoderms. Paleobiology 33(1):149-163]. To the list of references concerning Fell's identification of Kirk's ophiuroid as Ophiomyxa (p. 415), add that his first mention was in his 1960 key to the genera of ophiuroids. Additional and more recent sources of information relating to Table 2 include: Emlet, R.B. 2006. Direct development of the brittle star Amphiodia occidentalis (Echinodermata,Ophiuroidea, Amphiuridae) from the northeastern Pacific Ocean. Invertebrate Biology 125(2): 154-171 (confirms table entry regarding place of hydrocoele closure and location of the stone canal). ALSO Morris, V.B. 2011. Coelomogenesis during the abbreviated development of the echinod Heliocidaris erythrogramma and the developmental origin of the echinoderm pentameral body plan. Evolution & Development 13(4):370-381 (confirms table entries for echinoids and asteroids regarding place of hydrocoel closure and location of stone canal). Further to Table 3, Blastoids entry, refer also to Horowitz, Able & Strimple, 1986, Abnormalities in Pentremites ...etc., Journal of Paleontology 60(2):390-399. Also further to Table 3, NEW ENTRY FOR HOLOTHURIANS: from plate 1A of Semon (1888) the relation of the larva to the imago is traced through sequential diagrams; these diagrams show that the larval anus is carried through to the adult and lies ventrally in the plane of hydrocoel closure; this places the larval axis/plane coincident with the A-CD axis/plane of edrioasteeroids; displacement of the anus to the distal end of the holothurian where it cannot be assigned to an interradeal position is subsequent; the reference is Semon, R., 1888, Die Entwickelung der Synapta digitata und die Stammesgeschichte der Echinodermen, Jenaischen Zeitschrift fur Naturwissenschaft 22(N.F. XV): 135 pp., 7 plates. Further to Note 2 discussion of alternating versus opposite ambulacral plates: Stenaster has opposite ambulacral plates, but in one arm of specimen MPRI 0109 of Stenaster obtusus the ambulacrals are definitely alternating and are convex zig-zag. The place where the alternation starts has not been discerned, but this specimen shows that newly formed plates (formed by the ocular plate rule) maintain this aberrant alternating pattern, indicating that the pattern is an epigenetic response to a template. The reasoning begins from realizing that a local defect will cause local accomodation, it will not cause a complete series of plates to shift register. The shift of register must have been present and perpetuated at the time that the plates were laid down at the growing tip of the arm. This means that once the template switches from an ambs-opposite template to an ambs-alternating template, the alternating pattern is maintained. One aspect of significance to this observation is that it calls into question the appraisal and meaning to attach to examples described as ambs-opposite proximally but ambs-alternating distally. Specimen MPRI 0231A is a protasterid ophiuroid (cf. Bohemura) that has alternating ambulacral plates except that there is a place in one arm that has three successive pairs of ambulacrals that are aberrantly opposite. Further to evidence on changes of symmetry axes (Note 5): In the Order Echinoida, the oval test of Parasalenia gratiosa var. boninensis is elongate in the IIIb-5b direction, that of Echinometra mathaei in the 3a-Ia direction, that of Hetrocentrotus mamillatus and of H. trigonarius in the IVb-1b direction, and the test of Colobocentrotus mertensii is elongate in the 4a-1b direction using Lovén's nomenclature (Hayato Ikeda, 1939, The shape of the test of Colobocentrotus mertensii Brandt (Echinoidea, Echinometridae), Annotationes Zoologicae Japonenses 18(3):194-201, pls. 9-10). A specimen of C. mertensii with its long and short axes apparently interchanged, as judged by the location of the madreporite, may be a case of situs inversus (unpublished suggestion of FHCH; specimen described by Hiroshi Ohshima, 1939, Some rare abnormalities in asteroids and echinoids,Zool. Mag. Tokyo, 51:158-162). The ovoid test of Tiarechinus can also be mentioned (Treatise on Invertebrate Paleontology 1966 Part U pp. U436-U437). The test is circular but the apical system is elongate in the III-5 direction in Orthopsis ruppellii, Order Orthopsida (Treatise fig. 326,1h), likewise in Glyphocyphus (G.) radiatus, Order Temnopleuroida (Treatise pp. U414-415). Further to evidence of the axis of the bilateral larva in the adult (Note 7): The manner in which J.E. Smith assigned Roman numerals to the rays of starfish is explained by Smith in the 1966 book Physiology of Echinodermata (R. A. Boolootian, editor; see page 508 figure caption: madreporite between rays I and V; this is the same system as Ludwig 1899, also Kjerschow-Agersborg 1922, also Rodenhouse & Guberlet 1946). Translated into my 1998 table of ray homologies, the most frequently leading ray in Asterias according to Smith is ray C, and the associated axis for this bilateral tendency is the C-EA axis. [As an aside, it surprises me that Smith assigned numbers in the manner of Ludwig 1899 and not in the manner of Chadwick (1923 LMBC Memoire on Asterias). His leading ray observations would be more consistent with the reports for other starfish if he was using the Chadwick/Gemmill system.] The details for SET I specimen numbers 106-112 are listed below: 106. Protasteridae, Eugasterella africana Jell & Theron, South African Museum, Cape Town, paratype, SAM K1015, figured Jell & Theron (1999:177, Fig. 48A), madreporite in III/IV interradius. 107. Protasteridae, Eugasterella africana Jell & Theron, South African Museum, Cape Town, paratype, SAM K1014, figured Jell & Theron (1999:177, Fig. 48C) , madreporite in III/IV interradius. 108. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561B, paratype, figured Jell & Theron (1999:177, Fig. 48B), madreporite in III/IV interradius. 109. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561D, paratype, figured Jell & Theron (1999:176, Fig. 47C), madreporite in III/IV interradius. 110. Protasteridae, Eugasterella africana Jell & Theron, Geological Survey of South Africa, Bellville in Capetown No. B4561a, holotype, figured Jell & Theron (1999:175, Fig. 46A), madreporite in III/IV interradius. 111. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B0196, figured Jell & Theron (1999:181, Fig. 51C), madreporite in III/IV interradius. 112. Cheiropterasteridae, Hexuraster weitzi (Spencer), Roy Oosthuizen Collection, Zwartskraal, Prince Allbert No. RO 45, figured Jell & Theron (1999:163, Fig. 39) , madreporite in III/IV interradius. The details for SET II specimen numbers 215-219 are listed below: 215. Protasteridae sp. Specimen not figured by Jell & Theron. Specimen details still not known. Arms score ?AB*A? 216. Eophiuridae, Haughtonaster reedi Rilett, Geological Survey of South Africa, Bellville in Capetown No. B4567, figured Jell & Theron (1999:158, Fig. 36B). Arms score AA?*AB 217. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4512, figured Jell & Theron (1999:183, Fig. 53). Arms score AAB*?? 218. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4513, figured Jell & Theron (1999:182, Fig. 52D). Arms score ?AB*?B 219. Protasteridae, Strataster ohioensis Kesling & LeVasseur, , Geological Survey of South Africa, Bellville in Capetown No. B4513, figured Jell & Theron (1999:179, Fig. 49b). Arms score AAB*A? NEW OBSERVATION: Specimen SUG299 of Encrinaster tischbeinianus (Roemer), Family Encrinasteridae, from De Doorns, Ceres District, Lower Devonian Bokkeveld Group, property of the Geological Collections, Stellenbosch University, South Africa, oral surface, figured Jell & Theron (1999:164, Fig. 40B-D): The specimen has a madreporite (enlarged in their Fig. 40D; madreporite not recognized by FHCH in the contact print in 1994). Using a magnifier to examine their Fig. 40C, the arms score AAB*AB. The five arms conform with Lovén’s AABAB rule. The madreporite is located in the III/IV interradius. The specimen fulfills SET I criteria and is assigned Set I reference number 113. Including the new observation, Lovén’s rule applies to the ophiuroid families Eophiuridae, Cheiropterasteridae, Encrinasteridae and Protasteridae (not just the Protasteridae as originally reported). The arms of specimen MPRI 0105 of Bundenbachia beneckei (Hunsruck Slate) score as AABAB, but the madreporite is not observable.

Hotchkiss, F. H. C. 1993. A new Devonian ophiuroid (Echinodermata: Oegophiurida) from New York State and its bearing on the origin of ophiuroid upper arm plates. Proceedings of the Biological Society of Washington, vol. 106, no. 1, pp. 63 84. Further to the comment on p. 73 regarding the very straight arms of the holotype and the existence of a stiffening reaction in ophiuroids, see remarks of W.K. Spencer (Monograph p. 246 citing von Uexkull, and p. 316 on Furcaster with arms stretched out very stiffly) that a very strong stimulus to the arm produces a rigor ... and the arms feel like small sticks. On the subject p. 75 that upper arm plates do not stay with the arm segments that become included in the disk, Ishida & Inoue (1993:104 Ophiura sarsii) noted that "about 4 dorsal arm plates at the base of the arm are incorporated into the disk". Further to the evidence p. 76 against the idea that upper arm plates get incorporated into the radial shields, note A.M. Clark 1966 Echinodermata [Port Phillip Survey 1957-1963], Memoirs of the National Museum, Melbourne No. 27, pp. 289-384, on p. 337 regarding Amphiura (Ophiopeltis) parviuscutata that basal arm segments have their dorsal arm plates reduced or even absent inside the disk area. Regarding the possible origins and homologies of the radial shields, these were not treated comprehensively on pp. 75-76 except where there was some suggestion that related radial shields to upper arm plates. For other origins of radial shields see for example Spencer (Monograph p. 244) and Matsumoto (Monograph p. 369). The statement of a reviewer that "it is incorrect to infer from Lyman's illustrations that radial shields grow by adding platelets" is true only for ophiurid ophiuroids in which radial shields grow from a single tiny scale which initially formed at the edge of the disk. Each radial shield of Amphioplus abditus originates from a single element (emphatically not by fusion of scales; Hendler 1978 p. 88). On the other hand, Brian Stewart showed that the radial shields of Astrobrachion constrictum grow by proximal sequential addition of scales of stereom to the proximal ends (1966. Growth dynamics of the radial shields of the euryalid snake star Astrobrachion constrictum. Invertebrate Biology 115(4):321-330). This confirms Fell's (1963:420) description of euryalid radial shield development. On the subject of retaining for the moment the two suborders Lysophiurina and Zeugophiurina p. 80, note W.K. Spencer (Monograph p. 9) on " remarkable analogous (homoplastic) course of development" of two series, and that "For the present both series are included in the Ophiuroidea". I do not regard Strataster as ancestral to crown group ophiuroids, but Smith, Paterson & Lafay (1995) depict it as ancestral in their cladogram. Contrary to my views, the cladogram of Juliette Dean Shackelton (2005 J. systematic Palaeontology 3:29-114, her fig. 15) does not maintain the integrity of the Lysophiurina/Zeugophiurina lineages. In particular, I do not accept her revision of the family Protasteridae to include both lysophiurine and zeugophiurine genera. It is my working hypothesis that her cladogram erroneously brings together ophiuroids that are in separately evolving lineages; it is my working hypothesis that all instances of Loven's law are plesimorphic and that none are homoplasic. A paper by Turner & Heyman, 1995, Proc. Biol. Soc. Wash 108(2):292-297 has important observations on the presence of radial shields and dorsal arm plates in Ophiosyzygus and many other ophiomyxids.

Hotchkiss, F.H.C., S.E. Churchill, R.G. Gelormini, W.R. Hepp, R.J. Rentler & M.T. Tummarello. 1990. Events of autotomy in the starfish Asterias forbesi and A. vulgaris. Presented at 7th International Echinoderms Conference, Sept. 9/14, 1990, Atami, Japan. Published 1991, pp. 537-541 in T. Yanagisawa et al. (eds.), Biology of Echinodermata, A.A. Balkema, Rotterdam, 590 pp. Prepublication results were cited by Emson & Wilkie (1980 Fission and autotomy in echinoderms. Oceanogr. Mar. Biol. Ann. Rev. 18:155-250; p. 222). CORRECTION: Our identification of the tourniquet muscle based on Hotchkiss (1979) had errors of interpretation that were corrected by Wilckie et al. [I.C. Wilkie, G.V.R. Griffiths & S.F. Glennie 1990. Morphological and physiological aspects of the autotomy plane in the aboral integument of Asterias rubens L. (Echinodermata). In: C. DeRidder, P. Dubois, M.C. LaHaye, M. Jangoux (eds.), Echinoderm Research. Rotterdam: Balkema, pp. 301-313; see p. 305 histology of the transverse basal ridge], but their paper was not yet published by the time of the Atami IEC. This does not negate our observations on the locus of the autotomy plane or on recognizing slow versus fast aspects of autotomy. In the meanwhile there continue to be new significant papers on autotomy in Asteroidea. ADDITIONAL PAPERS ON AUTOTOMY IN ASTEROIDEA: [J. M. Lawrence. 1992. Arm loss and regeneration in Asteroidea (Echinodermata). Pp. 39-52 in L. Scalera-Liaci and C. Canicatti (eds.) Echinoderm Research 1991. Rotterdam: Balkema.] [J. Marrs, I.C. Wilkie, M. Skold, W.M. Maclaren, and J.D. McKenzie. 2000. Size-related aspects of arm damage, tissue mechanics, and autotomy in the starfish Asterias rubens. Marine Biology 137:59-7.] [I. C. Wilkie. 2001. Autotomy as a prelude to regeneration in echinoderms. Microscopy Research Technique 55:369-396.] [I. C. Wilkie. 2010 Do gulls benefit from the starfish autotomy response? Marine Biodiversity Records (2010), 3 : e12 Cambridge University Press doi:10.1017/S1755267209990480.] [John M. Lawrence and Carlos F. Gaymer. 2012. Autotomy of rays of Heliaster helianthus (Asteroidea: Echinodermata). Zoosymposia 7:173-175.]

Hotchkiss, F. H. C. 1982. Ophiuroidea (Echinodermata) from Carrie Bow Cay, Belize. Pp. 387-412 in The Atlantic Barrier Reef Ecosystem at Carrie Bow Cay, Belize, I. Structure and Communities. K. Rutzler and I.G. Macintyre, editors. Smithsonian Contributions to the Marine Sciences, No. 12.

Corrections to identifications in this paper were published by Hendler & Peck (1998. Ophiuroids off the deep end: Fauna of the Belizean fore-reef slope. Pp. 411-419 in Echinoderm Biology, Burke et al. [eds.], Balkema, Rotterdam), as follows: Ophiomitrella glabra (H.L. Clark) is regarded as a junior synonym of Ophioblenna antillensis, Axiognathus squamatus (Delle Chiaje) as a junior synonym of Amphipholis squamata, and the six-rayed fissiparous Ophiostigma sp. was nominally referred to Ophiostigma isocanthum [but see below]. The Ophioderma sp. juvenile is Ophioderma rubicundum, and the Sigsbeia murrhina Lyman sensu Hotchkiss specimens are Sigsbeia conifera Koehler. The Ophiurochaeta littoralis Koehler sensu Hotchkiss specimens were reidentified as an undescribed species of Ophiurochaeta [four undescribed Ophiurochaeta species provisionally designated A, B, C and D are mentioned by Hendler and Peck]. In a subsequent paper the six-rayed fissiparous Ophiostigma sp. was described as the new species Ophiostigma siva by Hendler [1995. New species of brittle stars from the Western Atlantic, Ophionereis vittata, Amphioplus sepultus, and Ophiostigma siva, and the designation of a neotype for Ophiostigma isocanthum (Say) (Echinodermata: Ophiuroidea). Contributions in Science, Natural History Museum of Los Angeles County No. 458].

Hotchkiss, F.H.C. 1980. The early growth stage of a Devonian ophiuroid and its bearing on echinoderm phylogeny. Journal of Natural History 14:91 96. The paper documented primary rosette plates in an oegophiurid with alternating ambulacral plates (Lysophiurina). The Lysophiurina and Zeugophiurina lineages diverged no later than Early Ordovician time. The Zeugophiurina are the presumed stem group of the Silurian-Recent Ophiurida. Documentation of a primary rosette in Paleozoic Ophiurida can be mentioned: in Ophiaulax decheni (Devonian, Ophiurinidae) [R. Haude & E.T. Thomas. 1983. Ophiuren (Echinodermata) des hohen Oberdevons im nördlichen Rheinischen Schiefergebirge. Paläont. Z. 57(1/2):121-142]; in Aganaster gregarius (Mississippian, Ophiolepididae) [F.H.C. Hotchkiss & R. Haude. 2004. Observations on Aganaster gregarius and Stephanoura belgica. Pp. 425-431 in Heinzeller & Nebelsick (eds.), Echinoderms: München]. Additional references on postlarval growth stages in Recent ophiuroids include: -- Sumida, P.G. and P.A. Tyler. 1998. Postlarval development in shallow and deep-sea ophiuroids (Echinodermata: Ophiuroidea) of the NE Atlantic Ocean. Zoological Journal of the Linnean Society 124:267-300. -- Stöhr, S. 2005. Who's who among baby brittle stars (Echinodermata: Ophiuroidea): postmetamorphic development of some North Atlantic forms. Zoological Journal of the Linnean Society 143:543-576 [includes information on the euryalid Asteronyx loveni: the plates of the primary rosette are conspicuous at first, but do not endure: at 3.2 mm disk diameter they are small fragments at the center of the disk]. -- Stöhr, S., and M. Segonzac. 2006. Two new genera and species of ophiuroid (Echinodermata) from hydrothermal vents in the east Pacific. Species Diversity 11(1):7-32 [p. 26 and fig. 7A, ophiacanthid Ophiolamina eprae postlarva 0.7 mm disc diameter has "six instead of the usual five radial primary plates, which is probably an aberration"]. This aberration resembles Hamling's ophiuroid rosette.

Hotchkiss, F. H. C. 1979. Case studies in the teratology of starfish. Proceedings of The Academy of Natural Sciences of Philadelphia 131:139-157. The historical review on pp. 139-143 is only a sampling of what could have been included. Some additional references, in historical order, follow: 1882. Bell, F.J. Note on the echinoderm fauna of the Island of Ceylon, together with some obversations on heteractinism. Ann. Mag. Nat. Hist., ser. 5, 10:218-225; 1887. Bell, F.J. The echinoderm fauna of the island of Ceylon. The Scientific Transactions of the Royal Dublin Society, series 2, 3:643-657, pls. 39-40. [an oblique cut regenerates obliquely to the arm axis]; 1900. Bedford, F.P. On echinoderms from Singapore and Malacca. Proceedings of the Zoological Society of London, pp. 271-299 [Archaster typicus with constriction at the base of one arm accompanied by superomarginals meeting across the abactinal surface, but the actinal plates are unaffected]; 1912. Richters, C. Zur Kenntnis der Regenerationsvorgange bei Linckia. Zeitschrift fur Wissenschaftliche Zoologie 100:116-175; 1914. Schapiro, J. Uber die Regenerationserscheinungen verscheidener Seesternarten. Arch. f. Entwmeck. 38:210-251; 1915. Nusbaum, J. & M. Oxner. Zur Restitution bei dem Seestern Echinaster sepositus Lam. Zool. Anzeiger 46(6):161-167; 1917. Zirpolo, G. Notizia di alcuni Asteroidi anomali pescati nel Golfo di Napoli, Echinaster sepositus Gray e Asterias glacialis O.F. Muller. Bollettino della Societa di Naturalisti in Napoli 30:20-29; 1918. Zirpolo, G. Un caso di rigenerazione parziale delle braccia in un Astropecten aurantiacus L. Pubblicacazioni della Stazione Zoologica di Napoli 2(2):169-175, pl. 10 [a 5-rayed individual loses two adjacent rays and some disk; only a single ray regenerates, resulting in a 4-rayed individual]; 1924. Zirpolo, G. Ulteriori notizie di Asteroidi anomali. Bollettino della Societa di Naturalisti in Napoli 36:305-346, pls. 6-8; Zirpolo, G. Notizia di un Echinaster sepositus Gray con sei braccia nel Golfo di Napoli. Atti Pontif. Accad. Nuovi Lincei, Anno 77:161-163. [No external or internal irregularities, providing his argument for hexamery from the first moment of life]; 1926. O'Donohue, C.H. On the summer migration of certain starfish in Departure Bay, B.C., Canada. Fisheries Research Board of Canada. Contributions to Canadian Biology and Fisheries 1:455-472; Zirpolo, G. Gemmazioni, rigenerazioni ipertipiche ed ipotipie studiate nell'Astropecten aurantiacus L. Bollettino della Societa di Naturalisti in Napoli 38:167-221 [includes examples of tetramery arising when two arms regenerated in place of three, and when one arm regenerated in place of two]; 1927. Rose, M. & R. Dieuzeide. Sur un Astropecten aurantiacus L. anormal. Bulletin de la Société d’Histoire Naturelle de lAfrique du Nord 18(6):135-137; 1928. Zirpolo, G. 1928. Caso di eteromorfosi in un Astropecten aurantiacus L. Bollettino della Societŕ dei Naturalisti in Napoli. 39(ser. 2, vol. 19): 195-206 + pl. 17 [regeneration/development of the aboral skeleton and the marginal plates of the arm without there being an adjacency of the aboral side of the regenerating ray with the aboral disk]; 1929. Zirpolo, G. Le forme cometoidi dell'Asterias tenuispina Lmk. Bollettino della Societa di Naturalisti in Napoli 40:25-35; Zirpolo, G. Nuovo caso di gemmazione in un Astropecten aurantiacus L. Bollettino della Societa di Naturalisti in Napoli 40:83-87; Zirpolo, G. Notizia di Asteroidi irregolari. Bollettino della Societa di Naturalisti in Napoli 40:221-231; 1931. Abeloos, M. Les potentialites regeneretrices de la face dorsal des bras des Asteries, Trifurcation dorsal d'un bras chez Solaster papposus (Linck). Bull. Biologique de la France et de la Belgique fonde par Alfred Giard, 65(3):394-405. 1933. Mortensen, Th. The Godthaab Expedition, 1928, Echinoderms. Meddelelser om Gronland 79(2):1-62 + plate I. [on p. 17 arm number in Solaster papposus: nearly all have 10 arms, one has 12, a few have 9, one has 8 arms]; 1938. Domantay, J.S. An unusual bud due to heteromorphosis in Echinaster luzonicus (Gray). Phillipp. J. Sci. Manila 64:281-283; 1941. Fisher, W.K. A new genus of sea stars (Plazaster) from Japan, with a note on the genus Parasterina. Proc. USNM 90(3114):447-456, pls. 66-70; 1945. Fisher, W.K. Unusual abnormalities in sea-stars. J. Wash. Acad. Sci. 35:296-298; 1948. Tortonese, E. Variazioni fenotipiche e biologia della populazione di Astropecten aranciacus (Echinodermi) del Golfo di Napoli, con riferimenti a specie congeneri. Bollettino dell'Istituto e Museo di Zoologia dell'Universita di Torino 1(9):87-123 [fig. 2 of arm developing beneath disk]; 1951. Alfano, B. Commemorazione del Prof. Giuseppe Zirpolo. Bollettino della Societa di Naturalisti in Napoli 59:39-64 [on the life and publications of G. Zirpolo]; 1958. Dollfus, R.Ph. Courbe interbrachiale chez une asterie du genre Stellasteropsis. Bull. Soc. Zool. Fr. 83:294-297 [compares 4-armed and 5-armed individuals]; 1967. Raup, D.M. & E.F. Swan. Crystal orientation in the apical plates of abberant echinoids. The Biological Bulletin 133(3):618-629 [ocular IV and genital 4 are judged missing in two four-part urchins; of special interest because interradius IV/V is the location of closure of the hydrocoele crescent to form the ring canal; ray IV is on the ventral or posterior horn of the hydrocoele crescent]; 1969. Kenny, R. Growth and asexual reproduction in the starfish Nepanthia belcheri (Perrier). Pacific Science 23(1):51-55. 1975. Hinegardner, R.T. Morphology and genetics of sea urchin development. American Zoologist 15:679-689 [breeding experiments and loss of symmetry control]; 1976. Ebert, T.A. Natural history notes on two Indian Ocean starfishes in Seychelles: Protoreaster lincki (de Blainville) and Pentaceraster horridus (Gray). J. mar. biol. Ass. India 18(1):71-77 [most of the atypical arm numbers are probably due to damage during early post-metamorphosis]; 1986. Horowitz, A., S. Able, & H.L. Strimple. Abnormalities in Pentremites Say (Blastoidea) from the Pella Formation (Upper Mississippian) of Iowa. Journ. Paleontology 60(2):390-399; 1989. Ausich, W.I. & T.W. Kammer. Teratological specimen of Agaricocrinus americanus (Roemer) (Lower Mississippian, Crinoidea). J. Paleont 63(6):945-946; 1991. Boursin, J. Un curieux (b)oursin ou coussin? Xenophora, number 53:25 [square 4-armed Halityle regularis from New Caledonia]; 1992. Ramalingam, K. & A. J. Antony. A note on the tetramerous starfish (Oreaster sp.). Comparative Physiology and Ecology 17(3):121-122; 1999. James, D.B. Abnormal asteroids from the seas around India. Mar. Fish. Infor. Ser., T. & E. Ser., number 158:21-22; 2000. Hotchkiss, F.H.C. On the number of rays in starfish. American Zoologist 40;340-354 [this paper inludes additional references not repeated here, as well as a new synthesis for the topic]; 2002. Dupont, S. & J. Mallefet. Abnormal forms in the brittle-star Amphipholis squamata: a field study. J. Mar. Biol. Ass. U.K. 82:491-493 [deviations from pentamerism are not a heritable character but are a consequence of environmental perturbations on the metamorphosis of larvae and/or abnormal regeneration of arms]; 2004. Yuriy Mitrofanov and O.V. Dogadova. Disorder of symmetry in the marine starfishes of Amursky Bay. pp. 177-178 in Proceedings of the Thrd Workshop on the Okhotsk Sea and adjacent areas. PICES Scientific Report No. 26; 2005. Chessa, L.A., F.P. Patti, and G. Delrio. Peltaster placenta (Muller & Troschel 1842) forma tetramera: un ritrovamento insolito. Biologia Marina Mediterranea 12(1):253-256; 2007. Ceranka, T. Symmetry disorders of the test of the Miocene echinoid Echinocyamus from Poland. Acta Palaeontologica Polonica 52(3):503-518; 2009. Sigovini, M., and D. Tagliapietra. Segnalazione di un esemplare esaraggiato di Asterina gibbosa (Pennant, 1777) in Laguna di Venezia (Echinodermata: Asteroidea). Bull. Mus. civ. St. nat. Venezia 59:69-74; 2012. Prabhu, K., and S. Bragadeeswaran. Occurrence of abnormal starfish Astropecten indicus (Doderlain, 1888) (Echinodermata: Asteroidea) along southeast coast of India. Revista Biotemas 25(4):293-296; 2013. Chamundeeswari, K., S. Saranya, S. Shanker, D. Varadharajan, and S. Rajagopal. New occurrence of abnormal sea star, Astropecten indicus from Mudasalodai, south east coast of India. Cell & Developmental Biology 2:116. doii:10.4172/2168-9296.1000116. 2014. S. Shanker and P. Vijayanand. Abnormal starfish, Pentaceraster regulus from Thondi, East Coast of India. Cell & Developmental Biology 3:135. doi 10.4172/2168-9296.1000135. 2015. G. Chelladurai, S. Balakrishnan, G. Jayanthi, K.K. Ajeesh Kumar and J. Mohanraj. Report on the occurrence of abnormal four-armed red-knobbed starfish Protoreaster linckii (Echinodermata: Astroidea), Tuticorin coast, south-east coast of India. Marine Biodiversity Records, Volume 8.

Behavioral evidence for a physiological anterior in locomotion in starfish was examined on pp. 153-154. Concerning behavioral evidence for nonrandom use of arms in ophiuroids see: Welte, N.T. & L.M. Lutton. 2003. Appendage selection during brittlestar locomotion. Journal of the Pennsylvania Academy of Science 77(1):15-19. This paper revealed a pattern of two adjacent arms being used least for leading and most for propulsion. The identity of the arms with respect to the location of the madreporite was not examined and deserves further study. See also Parker, G.H. 1936. Direction and means of locomotion in the regular sea-urchin Lytechinus. Musée Royal d’Histoire Naturelle de Belgique, Bruxelles, Memoire series 2, number 3, pp. 197-208. See also Roderick MacDonald. 1936. A study of symmetry in the Centrechinoidea, based on behavior, with special reference to Lytechinus variegatus. Proceedings of the American Philosophical Society 76(1):87-123. See also Yoshimura, K. & T. Motokawa. 2008. Bilateral symmetry and locomotion: do elliptical regular sea urchins proceed along their longer body axis? Marine Biology 154:911-918. See also Yoshimura, K. & T. Motokawa. 2009. In which direction do regular sea urchins walk? [abstract] IEC Meeting, Hobart, Tasmania, January 2009, book of abstracts, page 86. See also Yoshimura, K., and T. Motokawa. 2010. Bilaterality in the regular sea urchin Anthocidaris crassispina is related to efficient defense not to efficient locomotion. Marine Biology 157:2475-2488. See also Yoshimura, K., T. Iketani, and T. Motokawa. 2012. Do regular sea urchins show preference in which part of the body they orient forward in their walk? Marine Biology 159:959-965. Concerning Henricia in Massachusetts: "The Cribrella moves usually with two of the arms turned backward, and the three others advanced together, the two posterior ones being sometimes brought so close to each other as to touch or their whole length." The accompanying drawing seems to show the madreporite between the two appressed rays [source pp. 112-113, Agassiz, E.C. and A. Agassiz, 1865, Seaside studies in natural history, marine animals of Massachusetts Bay, Radiates; Ticknor and Fields, Boston, 155 pp.]. See also Jones, D.A., E.W. Knight-Jones, J. Moyse, P.C. Babbage, and A.R.D. Stebbing [1968. Some biological problems in the Aegean. Underwater Association Report, Malta 1968:73-78. -- leading arm in Echinaster sepositus on p. 77 is ray A of FHCH notation]

Hotchkiss, F.H.C. 1978. Studies on echinoderm ray homologies. Loven's law applies to Paleozoic ophiuroids. Journal of Paleontology 52(3):537-544.

Hotchkiss, F.H.C. 1977. Ophiuroid Ophiocanops (Echinodermata) not a living fossil. Journal of natural History 11: 377-380. Additional comments in Hotchkiss (1995): Ophiocanops does not have the 'auluroid' vertebrae of the Oegophiurida (p. 423); the extraordinary gonadal and stomachal characters perhaps are retained from the Oegophiurida; hence perhaps a surviving member of the stem group of the Order Phrynophiurida (p. 415, 423 and Fig. 6). Concerning the adoral plates filling the oral interradii and forcing the madreporite and the oral shields to be ambital: there is an instructive similar case of forcing the oral shields to the ambitus in juvenile specimens of Ophiozonella falklandica [stage with primary rosette and only three arm segments beyond the disc; source Mortensen 1936 Discovery Rpt. Echinoidea and Ophiuroidea, p. 303, fig. 31]. Am aware of the following 2001 paper but have not seen it: Fujita, T., S. Irimura & W.W. Kastoro. Biology of a rare ophiuroid Ophiocanops fugiens Koehler, 1922 (Echinodermata) associated with black corals, with notes on the specimens collected from Lembeh Strait, Bitung, Indonesia. In: Terazaki, M., A. Taira, M. Uematsu, Y. Michida and T. Kaneko, eds., Proceedings of the 11th JSPS Joint Seminar on Marine Science, Center for International Cooperation, Ocean Research Institute, University of Tokyo, Tokyo, 2001, pp. 326-333. For more on this topic see 2008 paper by Sabine Stöhr, Chantal Conand and Emilie Boissin: Brittle stars (Echinodermata: Ophiuroidea) from La Réunion and the systsematic position of Ophiocanops Koehler, 1922. Zoological Journal of the Linnean Society, 2008, 153:545-560 [they describe Ophiocanops multispina n.sp., transfer Renetheo felli to Ophiocanops, and present many new observations on morphology including upper arm plates, a second under arm plate, side arm plates that meet ventrally so that there is no resemblance to the ambulacral groove of Paleoozoic Oegophiurida, and place Ophiocanops in the Ophiomyxidae].

Hotchkiss, F. H. C. 1976. Devonian ophiuroids of New York State. Reclassification of Klasmura, Antiquaster, and Stenaster into the Suborder Scalarina nov., Order Stenurida.--New York State Museum Bulletin 425:1-39. In text-figure 1 the labels for the "cup for tube foot" and the "overlapping proximal and distal hinges" are wrongly attached. This error is corrected in Hotchkiss et al. 1999, J. Czech Geol. Soc. 44/3-4, Figure 1, page 330, corrected drawing. Additional Leintwardine specimens of Antiquaster magrumi: MCZ No. 385, BMNH 40300. Further to p. 10 studied material of Stenaster obtusus: 6-rayed E52410a,b, is the Gray Collection specimen D.38 mentioned by WK Spencer as 6-rayed (WKS Monograph p. 358 last sentence; Dr. Timothy A.M. Ewin, personal communication 16 October 2018).

some references to birds/turtles drumming for worms

American woodcock. " Later, at 10:30 a.m., this bird and another fed in bright sunshine on the lawn, nonchalantly pulling worms from the sod. They had found the only square footage in town that was frost free. Their side-to-side- shifting, accompanied by deliberate weight shifts from foot to foot, must cause enough vibration to stir the worms below." Joe Choiniere, director of Mass Audobon's Wachusett Meaadow Wildlife Sanctuary. Seasons of the woodcock: the secret life of a woodland shorebird. Sanctuary, The Journal of the Massachusetts Audubon Soociety, Summer 2006, 45(4):3-5. [see p. 5]

Gull in grass park beside Lake Merritt, Oakland, California, spring 1991. Observations by F. Hotchkiss communicated to Ned Newton. "... on the grass, to the front and left of me, only eight feet from the sidewalk, was a gull. The gull seemed to be stepping in place. I stopped to watch it on the grass. It stepped again... it did it clearly with an alternate, rapid step -- stepping in place. In between this stepping it would stop -- it would peer at the ground just in front of itself, but never walking -- just with its feet stationary, planted -- and I would then see it grab an earthworm with its beak -- and quickly swallow it -- and look for another -- and then resume the stepping in place -- without moving an inch. This gull was a talented earthworm drummer. How on earth did it ever learn to do this!!" Ned Newton added the following comments: "Well, it seems that I have heard of Robins and Woodcock drumming up worms. But it must have been fun to see a relatively large bird like a gull doing this. I was telling a friend from Connecticut about the drumming for worms. She was not at all surprised and added that she has seen turtles doing this same thing -- hopping from one foot to another. Somehow they don't have the speed that I would think is required. Anyone who has been out at night catching nightcrawlers knows that you have to be fast. Well, maybe the next time the kids are dancing it would be interesting to look around near by or on the flood and see if worms are raised to the surface." Newton's Notes, April 26, 1991 [Notes from the President], Nashoba Valley Bird Club, The Meadow Plover 2(2) Spring 1991.

On this topic see also an excellent entry on Wickipedia: European Herring Gull -- section on Diet -- reports drumming for worms

For information on turtles stomping for earthworms see:

Ernst, C. H., J. E. Lovich, and R. W. Barbour. 1994. Turtles of the United States and Canada. Washington, D.C.: Smithsonian Institution Press. [summary of information]

Kaufmann, J. H. 1986. Stomping for earthworms by wood turtles, Clemmys insculpta: a newly discovered foraging technique. Copeia 1986(4):1001-1004.

Kaufmann, J. H., J. H. Harding and K. N. Brewster. 1989. Worm stomping by wood turtles revisited. Bull. Chicago Herpetol. Soc. 24: 125-126.

Kirkpatrick, David T. and Catherine Kirkpatrick. 1996. Stomping for earthworms by Clemmys insculpta in captivity. Bulletin of the Chicago Herpetological Society 31(2):21-22.

sunflower sea star

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